Ernest Everett Just was an early twentieth century American experimental embryologist involved in research at the
Marine Biological Laboratory (MBL) at Woods Hole, Massachusetts, and the
Stazione Zoologica in Naples, Italy. Just was known for simple but elegant experiments that supported the
“fertilizing” theory of
Frank R. Lillie and served as an antagonist to Jacques Loeb’s work with
artificial parthenogenesis. Just’s many experiments with marine invertebrates showed that the egg surface, or
ectoplasm, plays an important role in the fertilization and development of eggs.

Just was born in Charleston, South Carolina, on 14 August 1883 to Charles Frazier Just Jr. and Mary Matthew Just. His father died in 1887 and soon after the Just family moved to James Island, off the coast of South Carolina. His early education consisted of attending the small school that his mother founded and directed. Just left James Island at the age of twelve to attend the Colored Normal Industrial Agricultural and Mechanics College at Orangeburg (now South Carolina State College). In 1899 Just graduated with a Licentiate of Instruction, meaning that he was certified to teach in any black school in South Carolina. He was only fifteen years old.

Teaching did not appeal to the teenager so he traveled north, picking up odd jobs along the way until he reached Kimball Union Academy in Meriden, New Hampshire. Just finished a classical course of study in three years, during which time his mother died. The young college graduate was advised by friends and teachers to attend Dartmouth College which he decided to do. At Dartmouth, Just studied biology, history, literature, and the classics. He graduated from Dartmouth in 1907, the only “magna cum laude” in his class, with an AB degree and joined the English faculty at
Howard University, Washington, DC in the fall of that same year. He was asked to take over the biology department and teach physiology in 1910, in addition to his English teaching duties. Soon after, Just became the first head of the new Department of Zoology and gave up teaching English courses.

Through a Dartmouth contact, Just communicated with Lillie at the University of Chicago about a post-graduate degree in biology. Lillie directed Just to begin research in 1909 at Woods Hole and to take courses at the
University of Chicago. Work at the marine station quickly turned into a research assistantship, working side-by-side with Lillie. All of this was done in addition to maintaining a full-time teaching position at Howard. After obtaining his PhD, Just returned annually to Woods Hole as an independent researcher.

Just focused his interests on marine invertebrate eggs, both in the laboratory and in their natural setting. Because of his tacit knowledge of how marine invertebrates reproduced in oceans and estuaries, Just was able to closely match his laboratory environment to that of the organism’s natural environment. In 1912, Just’s first paper,
“The Relation of the First Cleavage Plane to the Entrance Point of the Sperm.” was published in the
Biological Bulletin. Just showed that eggs of the marine worm
(Nereis) cleave in different planes depending on the sperm’s point of entry. To Just, the egg’s surface was an important and robust factor in the fertilization process. By showing that sperm had an equal probability of entering the egg at any point on the egg’s surface, and that the direction of cleavage depended on the arbitrary point of entry of sperm, and not some predetermined cleavage plane, Just made a dent in
preformationist theory. It was also during this time that he married Ethel Highwarden in 1912 and met
Jacques Loeb while Loeb was at the Rockefeller Institute for Medical Research. Similar research interests and Loeb’s stand on social equality made the two embryologists fast, but not longstanding friends. In 1915 Just was the first to receive the
Spingarn Medal, presented annually to the African-American who performs the greatest service to his or her race. This was followed by the University of Chicago awarding Just his PhD degree in experimental embryology in 1916.

During 1919 and 1920, Just published four papers in the Biological Bulletin, all focusing on his work with the
sand dollar Echinarachnius parma. In one set of experiments Just measured the elevation of the egg membrane at sperm contact and the time that it took for the membrane to be penetrated by a sperm. He observed that the sperm was pulled into the egg rather than the commonly held view that it actively bored its way into the egg. Just also documented a “wave of instability” that moved from the sperm’s entry point to the opposite side of the egg. Since then, embryologists have proven that such an instability wave is a wave of cortical granule exocytosis that forms the fertilization envelope. Just also saw that the wave was associated with an immediate blocking of any further sperm penetrability of the egg.

In 1920, while continuing to teach at Howard, Just obtained a ten-year research fellowship from
Julius Rosenwald through the National Research Council. He wasted no time in returning to his work at Woods Hole where he continued to study the process of fertilization with results that strengthened Lillie’s work and questioned Loeb’s idea of “superficial cytolysis.” According to Loeb’s studies, egg development could be initiated by exposing eggs to butyric acid. Development was then immediately followed by the release of lysine, Loeb’s cytolytic agent, to break down the egg cortex. Just showed that putting eggs in butyric acid for a short period of time actually slowed cytolysis rather than sped it up. Just was able to prove that the cytolytic effect of the butyric acid was due to overexposure of the eggs to the acid and nothing more. He went even further in dismissing Loeb’s experimental findings on
artificial parthenogenesis, attacking Loeb’s method of record-keeping and his apparent failure to maintain experimental conditions to mimic his experimental organisms’ natural environments.

The friendship that Just and Loeb had forged together at Woods Hole quickly vanished. Their disagreements played out for many years, with Loeb providing negative evaluations of Just to the Rockefeller Institute and the Carnegie Foundation. Every time Just tried to obtain grant money, Loeb’s evaluations seemed to rise up and quell any grant award.

In 1929 Just made his first trip to Europe and worked at
Anton Dohrn’s Stazione Zoologica in Naples. For six months he experimented with
sea urchins (Paracentrotus lividus and Echinus microtuberculatus) to see how these organisms develop and to continue testing Lillie’s “fertilizin” theory of fertilization. Around 1906 Lillie had hypothesized that eggs release a substance that he coined fertilizin. Upon contact with spermatozoa, said Lillie, fertilizin causes sperm to attach to it. Lillie believed that fertilizin molecules served as receptors on the egg’s surface. With receptors for egg and sperm surfaces, the molecule helped to “agglutinate” egg and sperm together.

In 1930 Just was invited to the Kaiser Wilhelm Institute in Berlin where he continued his studies of the ectoplasm with other species, including Amoeba. Just strove to show the importance of ectoplasm in initiation of development. While in Europe he published three articles on the role of ectoplasm:
“
Die Rolle des kortikalen Cytoplasmas bei vitalen Erscheinungen” (“The Role of Cortical Cytoplasmin Vital Phenomena”) in Naturwissenschaften (1931) and
“On Origin of Mutations” (1932) and
“Cortical Cytoplasm and Evolution” (1933), both published in the American Naturalist. All three articles pointed to Just’s view that ectoplasm is necessary for fertilization to occur.

With little hope of ever being able to teach anywhere but a black college, and his continued failure to secure research funds, Just went to Europe in 1938 with the intent of leaving Howard and finishing out his research career on a new continent. In 1939 he published two books:
Basic Methods for Experiments on Eggs of Marine Animals and
The Biology of the Cell Surface. Both books reflected Just’s holistic view of eggs and embryos: that is, eggs are to be taken seriously in their own right rather than seen simply as tools to manipulate in order to prove a theory. While Just’s experiments may have been simple, he was an intense perfectionist when it came to laboratory procedure. His ability to keep laboratory environments similar to actual marine environments aided in the integrity of his research. It also led to his life-long criticism of experimental embryologists who failed to appreciate his tacit knowledge about inducing marine invertebrate reproduction. To Just, too many embryologists were busy taking eggs out of natural environments and subjecting them to unnatural manipulations while ignoring the importance of the eggs’ environment as an important factor in development. In The Biology of the Cell Surface, Just also continued his attack on the role of genes in development. He remained adamant that cytoplasm was the key to development and not the nucleus. This countered the growing enthusiasm by geneticists who held the idea that the nucleus controlled fertilization and development.

The Nazi invasion of France in 1940 forced Just to return to the US and Howard University, one of the few institutions at the time that would hire a black scientist. His attempt to recareer again in the United States was short-lived however. Just died of pancreatic cancer on 27 October 1941.

Ernest Everett Just was an early twentieth century American experimental embryologist involved in research at the Marine Biological Laboratory (MBL) at Woods Hole, Massachusetts, and the Stazione Zoologica in Naples, Italy. Just was known for simple but elegant experiments that supported the "fertilizing" theory of Frank R. Lillie and served as an antagonist to Jacques Loeb's work with artificial parthenogenesis.

In 1888 when students and investigators arrived in
Woods Hole for the inaugural session of the
Marine Biological Laboratory (MBL), they recognized the need for a library collection of books and journals. The one wooden building on campus, later known as Old Main, housed everything, with researchers upstairs and the student laboratory downstairs. Lectures were held in one corner, and shelves held what books and journals were contributed. As the first MBL Director
Charles Otis Whitman noted in his 1888 Annual Report, having a library was absolutely essential for the success of the lab and would have to be provided somehow. The initial core volumes should include reference works and textbooks, and also the important journals in the four languages thought to be essential at the time.

By the second year,
Whitman’s report expressed gratitude for the many contributions to the library. For
Whitman, a “comprehensive biological library” would be the foundation on which a first rate laboratory would be built. Both research and instruction depend on such a resource. This early commitment to building a comprehensive journal collection as well as collecting the most important books has paid off, so that the library has been called a “national treasure” and has long been arguably the best complete and focused collection of life science journals available.

The vision for a great library was essential in attracting donations of funds to purchase journals and books, of course, but there were also other strategies for collecting. Visitors to the lab were invited to submit reprints and other research materials. When the
MBL began publishing
The Biological Bulletin in 1899, they immediately established an exchange program with other journals and publishers. This exchange program was critical especially in the years of WWI and again in WW II, when few libraries had funds to purchase volumes and international cooperation was more challenging. Yet it was possible to continue publishing copies of their own publications and to exchange them later when regular mailings resumed. As a result, the MBL-WHOI (the
Woods Hole Oceanographic Institution) library has complete runs of most journals even when other libraries are missing those difficult years. When funds were available, the
MBL purchased back issues to fill in incomplete runs and binding of individual issues into volumes became a priority very quickly (as mentioned in the fifth annual report). In 1895
Whitman urged that $1000 per year was needed just to sustain the current level of library acquisition.

In the report for the years 1896–1899,
Cornelia Clapp provided the first official “Report on the Library.”
Clapp had been the first student to arrive at the lab in 1888 and had returned as an investigator; she also became the first woman trustee in 1910 and served in that role until her death in 1934. It is fitting that she served as first librarian, enthusiastically growing and protecting the collection that she also used. Though referred to as “Miss Cornelia M. Clapp, Librarian,” she held a PhD from Syracuse University in 1888 and another PhD from the
University of Chicago, where she worked with
Whitman. In her first report she acknowledged the many gifts to the lab, including the accumulating files of papers contributed by lab researchers themselves. She appealed for more funds for purchasing and binding journals. This remained the theme for many years.

At first the library collection was housed in the shelves along one end of the wooden building. One year, many of the volumes seemed to have disappeared, but the next summer they were discovered tucked up into the roof, apparently for protection from storms though not successfully protected from all the birds. As the collections continued to grow, they added to the demand for more space. Finally, when the first permanent brick building was constructed in 1914, the library had a safe and protected home. This building, funded by second MBL Director
Frank Rattray Lillie’s father-in-law
Charles Crane, and named the Crane Building, gave the
MBL a way to demonstrate to all potential donors and supporters that the MBL intended to last forever and to make an impact with its research and teaching missions.

The library budget remained $1000 a year, and despite its new secure home, collections depended very much on donations. Individuals donated money and books, journal exchanges expanded, and the librarians persuaded publishers to donate volumes that were then put on a New Book shelf as advertising, so that visiting scientists would go back home and have their institutional libraries buy them. In 1913 H. Mc. E. Knower served as librarian and in his report strongly urged that the library needed an assistant to serve as a year-round librarian. Just having a volunteer scientist in the summer was not enough, since the collections were often left in a chaotic muddle of energetic use by the end of a season and there was nobody there during the rest of the year to straighten things out. Especially as the number of donated reprints grew, and as they received a great deal of use during the summer, it was considerable work just restoring the collection to order. Also, trying to keep on top of all the donations and exchanges during the summer alone was insufficient.

With a new building and library facility, it was time to hire a librarian. Miss
May E. Scott accepted the position and developed new catalogs, formally reaccessioned all the materials, and determined that the library had over 3300 volumes, plus about 1500 reprints. During the first year of her service, the library bound over five hundred volumes, replaced missing numbers, and added many more items. With a generous donation of over 2500 duplicates from the American Museum of Natural History, the library had achieved a new level of excellence.

Through the years, major donations have come at critical times from such groups and foundations as the
Carnegie Foundation, and the
General Education Board ($10,000 in 1926), so that after a period of intense growth, by 1926 the library had already grown to 18, 220 volumes plus a carefully catalogued 38,000 reprints.

In 1924 the library moved to what became the five permanent stacks in the Lillie Building—a substantial brick building that extended the Crane laboratories. The building was constructed with major donations of well over one million dollars, especially from
Rockefeller Foundation and
John D. Rockefeller, Jr. personally,
Carnegie Corporation, and
Charles Crane. The tremendous collaborative success shows just how highly the
MBL was regarded as a place of life science research and education.

After
Jane Fessenden became Librarian, the staff and collections grew considerably, as did their use. By the 1980s, it was becoming clear that the library was gathering a substantial collection, including some very valuable books and complete runs of journals that could not be replaced. The
Rare Books Room and Archives opened in the 1980s after
Cathy Norton took over as Librarian. The library moved to electronic publishing, with an emphasis on providing access for scientists in a way that successfully archives publications for continued use.

The
Rare Books Room and Archives contain rare books, of course, a catalog of which is available to anyone since the MBL-WHOI Library is committed to making materials available for use rather than preserving them in ways that exclude legitimate scholarly access. There are some artifacts, including a few items from courses or Albert Szent-Gyorgyi’s Nobel Prize for his work on vitamin C in Hungarian paprika peppers. The collection includes a few archival files, including some from
Frank Lillie that were transferred to the
MBL from the
University of Chicago, as well as some notebooks, scrapbooks, and letters. And the collection has brought together valuable historical research materials into library exhibits, including the
Leuckart Charts and other
collections. In addition, the
MBL is home to marvelous
MBL Library Photograph Collections, featuring early photographs dating back to before the
MBL was founded, a number of scrapbooks, and the wonderful
Alfred Frances Huettner Collection.

Today the library serves both the
MBL and
WHOI, based on a decision to combine resources to make an internationally leading library rather than to compete in the same small village of
Woods Hole. The MBL-WHOI library provides services for library researchers, some of whom draw mainly on the electronic journals, and Library Director
Cathy Norton has become a leader in promoting bioinformatics and extending the use of the collections through networks of users.

As a result, one might be tempted to think that there is no reason to come to the actual
MBL, since one can sit home and access modern journals on line. But this remains a vibrant place of science in the labs and in the courses. The library is a place where readers can find everything, pull it off the shelf, and see what else was going on in the same journal or at the same time. For at least the past decades, the MBL has seen a number of library readers who come precisely because they can find whatever they need “right there.” Recently, the library has added the formal category of Library Researcher, for those who come to spend a sabbatical, finish a major book project, or to collaborate with other scholars while using the library resources. The MBL-WHOI Library is very much an active place to find many kinds of wonderful materials but also a great place to find other people who know things and know where to find more materials. This place will never become obsolete because it is leading library information systems development, as through the
Encyclopedia of Life Project and the
Biodiversity Heritage Library. And this is also where the archival materials are housed and where scholars will find those materials and other scholars studying them.

In 1888 when students and investigators arrived in Woods Hole for the inaugural session of the Marine Biological Laboratory (MBL), they recognized the need for a library collection of books and journals. The one wooden building on campus, later known as Old Main, housed everything, with researchers upstairs and the student laboratory downstairs. Lectures were held in one corner, and shelves held what books and journals were contributed.

James David Ebert studied the developmental processes of chicks and of viruses in the US during the twentieth century. He also helped build and grow many research institutions, such as the Department of Embryology in the Carnegie Institution of Washington in Baltimore, Maryland and the Marine Biological Laboratory (MBL) in Woods Hole, Massachusetts. When few biologists studied the biochemistry of embryos, Ebert built programs and courses around the foci of biochemistry and genetics, especially with regards to embryology. He eventually directed the MBL's Embryology Course, and later, the MBL itself.

Ebert was born on 11 December 1921 in the town of Bentleyville, Pennsylvania. He attended public schools while growing up and then graduated from
Washington and Jefferson College in Washington, Pennsylvania in 1942. Not long after graduation he joined the United States Navy and eventually became a lieutenant. Ebert was stationed on a destroyer in the Pacific Ocean that was attacked by a kamikaze pilot. The destroyer sank and Ebert spent twenty-four hours in the ocean until being rescued. Afterwards, as a biologist, Ebert befriended and trained several Japanese developmental biologists.

In 1946 Ebert began working towards his PhD in developmental biology under the instruction of
Benjamin Willier at the
Johns Hopkins University in Baltimore, Maryland. In the same year he married Alma Goodwin, who was a Women Accepted for Volunteer Emergency during the war. Ebert received his PhD in 1950 and immediately became a member of the faculty at the
Massachusetts Institute of Technology (MIT) in Cambridge, Massachusetts. After one year at MIT, Ebert moved to
Indiana University in Bloomington, Indiana. Ebert became an associate professor of zoology by 1955, and he had started a program of experimental embryology. He studied
chick embryos and the processes by which the protein make-up of the embryos changed throughout development.

Six years after receiving his PhD, Ebert became the director of the
Carnegie Institution of Washington's Department of Embryology, in Baltimore, Maryland. Prior to Ebert's term as director, the department had had three other directors. The Institution's president, Caryl Haskins, had contemplated closing the department and starting something new. However, with some persuasion from
Willier, Haskins spoke with Ebert and decided to give him the opportunity to run the department. Ebert and Haskins agreed that the department needed to focus on the study of genes and their regulation as well as the ways cells influence one another. Haskins said that Ebert's youth and enthusiastic personality made Haskins believe that Ebert would provide a fresh perspective to the department.

Ebert argued that it was his job to recognize and to recruit new talent and then support them in their work. He stressed the use of biochemistry and genetics, which in the 1960s blended together to form molecular biology. During this time, Ebert started to study the relationship between muscle cell differentiation and the propensity to infection in the Rous sarcoma virus.

While still director of the
Carnegie embryology department, in 1970 Ebert also became the president and nonresident director of the
Marine Biological Laboratory (MBL) at Woods Hole. At the
MBL he researched, with Keiko Ozato, the response of murine lymphocytes to mitogens.

In 1977 Ebert ended his term at the
Carnegie Department of Embryology, but he remained the director of the
MBL. From 1978 until 1987, Ebert lived in Washington, D.C., and he was the president of the whole Carnegie Institution of Washington. As the institution's president, he made the decision to help build a large optical telescope in Chile at Las Campanas Observatory, and he worked towards the creation of a common campus for both Carnegie departments in Washington.

Ebert remained involved with scientific institutions for the rest of his life. When leaving one institution, he found another one to join. He retired from the Carnegie Institution in 1987 and became the president of the
Chesapeake Bay Institute at the Johns Hopkins University, where he was a professor of biology for six years. Ebert was elected to many societies including the
National Academy of Sciences, the
American Philosophical Society, the
American Academy of Arts and Sciences, and the
Institute of Medicine. He was the vice president of the
National Academy of Sciences from 1981 through 1993 and he also chaired its Government-University-Industry Research Roundtable from 1987 through 1993. His colleagues elected him as president of the
Society for the Study of Development and Growth, the
American Institute of Biological Sciences, and the
American Society of Zoologists.

In retirement, Ebert and his wife Alma spent half of each year in Woods Hole and at the MBL. Ebert and Alma died on 22 May 2001 in an automobile accident while en route to Woods Hole.

Sources

Ebert, James D. "An analysis of the effects of anti-organ sera on the development, in vitro, of the early chick blastoderm." Journal of Experimental Zoology 115 (1950): 351–77.

Ebert, James D. "An analysis of the synthesis and distribution of the contractile protein, myosin, in the development of the heart." Proceedings of the National Academy of Sciences 39 (1953): 333–44.

Ebert, James D. "The effects of chorioallantoic transplants of adult chicken tissues on homologous tissues of the host chick embryo." Proceedings of the National Academy of Sciences 40 (1954): 337–47.

Ebert, James D. "The formation of muscle and muscle-like elements in the chorioallantoic membrane following inoculation of a mixture of cardiac microsomes and Rous sarcoma virus." Journal of Experimental Zoology 142 (1959): 587–621.

James David Ebert studied the developmental processes of chicks and of viruses in the US during the twentieth century. He also helped build and grow many research institutions, such as the Department of Embryology in the Carnegie Institution of Washington in Baltimore, Maryland and the Marine Biological Laboratory (MBL) in Woods Hole, Massachusetts. When few biologists studied the biochemistry of embryos, Ebert built programs and courses around the foci of biochemistry and genetics, especially with regards to embryology.

Jane Maienschein is the daughter of
Joyce Kylander and
Fred Maienschein, and was born in Oak Ridge, Tennessee, on 23 September 1950. She attended MIT as a freshman and then transferred to
Yale University in 1969 when Yale decided to admit women undergraduates. In 1972 she graduated with an honors degree in History, the Arts, and Letters having written a thesis on the history of science. She then attended
Indiana University and studied with historian of embryology
Frederick B. Churchill, took courses with embryologist
Rudolf Raff, and learned how to do embryological laboratory research with
Robert Briggs. She received her MA in 1976 and a PhD in 1978, with a pre-doctoral Fellowship at the Smithsonian to study the history of microscopes and microscopy, and an NSF-funded dissertation improvement visit to the
Marine Biological Laboratory (MBL) to reproduce old embryological experiments and soak up the history and resources of the
MBL Library and labs. Maienschein’s scholarly research focuses on the history and philosophy of developmental biology.

In graduate school
Briggs helped Maienschein reproduce historical experiments using the dissertation study of
Ross Granville Harrison’s 1907 experiments on nerve fiber development.
Harrison had asked whether the neuroblast cell (which we would now call a neural stem cell) can reach out and develop its fiber by protoplasmic outgrowth or whether the cell required a pre-established bridge, as many of his contemporaries argued must be the case.
Harrison carried out the first ever tissue culture experiment, in which he got the neuroblast cells to grow when transplanted into an artificial medium of frog lymph.
Briggs and Maienschein discovered that carrying out the experiment with the techniques
Harrison described led to lots of nice bacterial and other unidentified cultures, but not nerve cells. Retracing
Harrison’s steps revealed that he had taken advantage of being temporarily housed near the bacteriologists at
Yale University and had used more sophisticated aseptic techniques than he described.

This work led Maienschein to an analysis of the role of the details of scientific practices and the value of carrying out “practical history,” as Edwin Clarke called it. She has also asked questions about the role of experiments in settling (or failing to settle) issues of theoretical debate. Her work in history of embryology has concentrated especially on the late nineteenth and into the twentieth century, including work done at the
Marine Biological Laboratory in Woods Hole, Massachusetts, and on issues of
morphogenesis and differentiation related to cell division. This research has led her to study stem cell research and regenerative medicine.

Maienschein is also a dedicated teacher who has received multiple awards, including the Arizona State University Parents Association Professor of the Year Chair, Regents’ Professorship, and President’s Professorship. In addition, she received the History of Science Society’s Joseph H. Hazen Education Prize Award. During the 105th United States Congressional session, in 1997 and 1998, she served as senior science advisor to Congressman Matt Salmon, who served on the Science Committee. She took a group of undergraduates to Washington, which led to their paper presentation at the 150th meeting of the American Association for the Advancement of Science, and that led to an invitation to write an editorial for Science. The students’ essay on “Scientific Literacy” remains the only publication in Science by undergraduates, and it led to a longer peer-reviewed article in Science Communication.

This personal exposure to the political context of science also led Maienschein to research reflecting more seriously on the social, political, and legal contexts of scientific research. Most productively, this has resulted in collaborative publications and projects with bioethicist
Jason Scott Robert and
Rachel Ankeny.

Maienschein served as the first president for the
International Society for History, Philosophy, and Social Studies of Biology (“Ishkabibble”) in 1989–1991, president of the
History of Science Society in 2008 and 2009, and in numerous other administrative rolls. She is Director of the
Embryo Project, along with
Manfred Laubichler.

Jane Maienschein is the daughter of Joyce Kylander and Fred Maienschein, and was born in Oak Ridge, Tennessee, on 23 September 1950. She attended MIT as a freshman and then transferred to Yale University in 1969 when Yale decided to admit women undergraduates. In 1972 she graduated with an honors degree in History, the Arts, and Letters having written a thesis on the history of science. She then attended Indiana University and studied with historian of embryology Frederick B.

Charles Otis Whitman was an extremely curious and driven researcher who was not content to limit himself to one field of expertise. Among the fields of study to which he made significant contributions were:
embryology;
morphology, or the form of living organisms and the relationships between their structures;
natural history; and behavior. Whitman served as director of several programs and institutions, including the Biology Department at the
University of Chicago, where he helped establish a new style of biology and influenced the work of many researchers of his generation, as well as future ones. He also served as first director of the
Marine Biological Laboratory (MBL) in Woods Hole, MA. Besides his considerable achievements with his own scientific research, Whitman was a tireless mentor who had many students who went on to achieve great success in the field of embryology.

Whitman was born in North Woodstock, Maine, to parents Marcia and Joseph Whitman on 14 December 1842. He grew up on a farm and developed an interest in natural history, particularly that of pigeons, at an early age. Whitman’s family was typical of the rural area where he grew up, and he was educated in the public school system, but despite his family’s lack of money he was highly motivated to receive a college education. Whitman earned money by teaching and tutoring in private schools, and in 1865 he began attending
Bowdoin College in Brunswick, Maine. Whitman was enrolled in the accelerated program and finished his degree in 3 years, graduating in 1868 with a BA. After graduation from Bowdoin College, Whitman took a position as Principal of Westford Academy in Massachusetts, where he remained for four years. He then moved to Boston to accept a position as instructor in natural science at English High School. This move was one of great importance, as it was in Boston that he became aware of Harvard University Professor of Zoology
Louis Agassiz and enrolled to become one of fifty participants in the first session of the summer marine biology program at the
Anderson School of Natural History on Penikese Island in 1873. This experience had a profound impact on Whitman as well as on other of
Agassiz’s students. In 1874 Whitman joined the Boston Society of Natural History and, after a second summer at Penikese, he decided to dedicate himself to the full-time study of zoology.

In 1875 Whitman went to study in Europe under
Anton Dohrn at the
Stazione Zoologica in Naples. After working with Dohrn in Naples, Whitman and his fellow Penikese Island student
Charles Sedgwick Minot moved to Leipzig, Germany. There, under the direction of parasitologist
Rudolf Leuckart, he learned the modern methods of embryology and
microscopy. Whitman received his PhD from the
University of Leipzig in 1878. His dissertation was
“The Embryology of Clepsine (glossiphonia)”, with an emphasis on the direct role of
cleavage in
histogenesis, or the differentiation of cells into specialized tissue and organs during growth. This research was instrumental in laying the groundwork for future studies of
cell lineage. Whitman found evidence that leech egg development was completely predetermined. This finding supported the
regulative theory of embryo development, according to which the whole embryo regulates the development of each cell, in contrast to the mosaic theory in which each cell develops independently, like a mosaic tile. His discoveries while working with the leech were instrumental to future taxonomical and morphological studies.

In 1879 he was offered a postdoctoral fellowship at the Johns Hopkins University but turned it down when he was invited to become Professor of Zoology at the Imperial University of Tokyo. He only spent two years there, but his short tenure was extremely influential. Eight of Whitman’s students there went on to become prominent zoologists, including four who held major chairs, affording him the informal title “father of zoology” in Japan. From November 1881 until May 1882, Whitman returned to the
Stazione Zoologica to study the
embryology, life history, and classification of the
dicyemids which led to the publication of a standard reference work on the parasite in 1883. From 1882 through 1886 Whitman worked as an assistant to
Alexander Agassiz at the
Museum of Comparative Zoology at
Harvard University. During this time Whitman also served as the editor for the Department of Microscopy at the
American Naturalist Magazine. After Harvard, Whitman took the job of tutoring amateur zoologist Edward Phelps Allis, Jr., in Milwaukee, Wisconsin. In addition to tutoring Allis, he took on the task of directing the very short lived
Allis Lake Laboratory. While there, Whitman oversaw the work of many researchers, including
William Morton Wheeler, who went on to become a prominent figure in the study of social insects.

During the summer of 1888 Whitman was invited to direct the newly established
Marine Biological Laboratory in Woods Hole, a position he held until 1908. In 1889 Whitman left the
Allis Lake facility to take the position of Chair of Zoology at
Clark University in Worcester, Massachusetts. In 1892 Whitman moved again to become head of the biology department at the newly founded
University of Chicago. There Whitman had several students who went on to make names for themselves in embryology. One of the most prominent was
Frank Rattray Lillie, who took over as director at the MBL after Whitman and succeeded Whitman at the
University of Chicago, as well. Whitman had many embryologist colleagues at Chicago, including Frank Rattray Lillie,
Jacques Loeb,
Franklin Paine Mall,
Albert Davis Mead,
Shosaburo Watase, and
William Morton Wheeler. Whitman remained at the University of Chicago until his death on 6 December 1910.

Whitman’s study of
sexual dimorphism, the morphological differences between male and female organisms of the same species, was an influence on
Oscar Riddle and his endocrinological research. Whitman’s 1898 paper
“Animal Behavior” contains many examples of innate, non-learned, behavior. In his later work, he analyzed the relation between innate and learned behavior and the ability of animals to adjust their behavior to new experiences. Whitman saw a similarity of variation in related species, and the trends of
evolutionary change in all species from the simplest of organisms to the most advanced. In 1900, when researchers were torn between the theories of mutation and selection, Whitman was a strong proponent of selection.

Whitman published papers and journal articles on every aspect of his work, but is probably best known for his posthumously published three-volume work
The Orthogenic Evolution in Pigeons, considered to be the first extensive study in comparative ethology. Whitman was instrumental in the founding of several journals and academic institutions, including the
Journal of Morphology, the
Biological Bulletin, and the
American Morphological Society which, through a merger with the Western Branch of the American Society of Naturalists (known as the Society of American Zoologists in 1901 and 1902), became the
American Society of Zoologists in 1902.

Whitman’s work significantly impacted the field of embryology. It greatly influenced the researchers of his generation as well as future generations. Whitman made significant contributions in the fields of embryology, morphology, taxonomy, and ethology. He published numerous books and papers in all of these subjects. Whitman was a mentor to biology students in several institutions around the world. Many of the institutions and publications he founded continue to be at the top of the field of embryology today.

Charles Otis Whitman was an extremely curious and driven researcher who was not content to limit himself to one field of expertise. Among the fields of study to which he made significant contributions were: embryology; morphology, or the form of living organisms and the relationships between their structures; natural history; and behavior.

Frank R. Lillie was born in Toronto, Canada, on 27 June 1870. His mother was Emily Ann Rattray and his father was George Waddell Little, an accountant and co-owner of a wholesale drug company. While in high school Lillie took up interests in entomology and paleontology but went to the University of Toronto with the aim of studying ministry. He slowly became disillusioned with this career choice and decided to major in the natural sciences. It was during his senior year that he developed his lifelong interest in embryology. Graduating with a BA in 1891 Lillie then moved to the
Marine Biological Laboratory (MBL) at Woods Hole, Massachusetts, to work and study with
Charles Otis Whitman, the founding director of the MBL. Lillie collected and studied
cell lineage side-by-side with some of the most prominent embryologists of the time:
Edmund B. Wilson,
Edwin G. Conklin, and
Aaron L. Treadwell. Along with his cell lineage studies, Whitman guided Lillie to work on the question of how blastomeres contributed to the formation of organs in fresh water clams.

In 1892 Lillie followed Whitman to the
University of Chicago zoology department where Whitman had accepted a chair appointment. In 1894 Lillie graduated with a PhD in zoology. His dissertation was a descriptive study of cell lineage in freshwater mussels. From 1894 through 1899 Lillie worked as an instructor at the
University of Michigan. In 1895 he married Frances Crane, sister of Chicago businessman Charles R. Crane. His brother-in-law would soon play a large role in introducing Lillie to the social elite in Chicago and helping to expand the MBL campus. During his marriage Lillie and his wife had five daughters, one son, and three adopted sons. Lillie briefly taught biology in upstate New York at Vassar College before returning to the University of Chicago as an assistant professor of embryology. In 1902 he was made an associate professor followed by a full professor in 1906. In 1908 Lillie published his classic book on chicken embryology,
Development of the Chick: An Introduction to Embryology. Along with writing the text, Lillie prepared a large series of serial sections of the
chick embryo at various stages to serve as illustrations. With revisions, the text and laboratory manual continue to be used to the present time, serving as one of the best accounts available on bird development.

In 1910 Lillie was made chair of the Department of Zoology. During this time he united the embryology program with the rest of the zoology department. With the combining of the departments, the budgets were also combined and Lillie was able to use his influence to obtain more money for embryological research. In 1931 Lillie was appointed dean of the Division of Biological Sciences at Chicago. After many years of distinguished teaching and research he was made the Andrew MacLeish Distinguished Service Professor of Embryology and the dean of biological sciences. In 1935 he was given emeritus professor status.

Although rarely written about, Lillie was a member of Chicago’s
Eugenics Education Society, a committee member of the Second International Eugenics Congress, and served on the advisory council for the Eugenics Committee of the United States. In the early 1920s Lillie envisioned an Institute of Genetic Biology that would gather data to examine
population problems, public health, and social control, but this never came to fruition.

Lillie is probably best known for his leadership at the Marine Biological Laboratory at Woods Hole. He organized the MBL’s first
course in embryology in 1893 and became course director the following year. At that time the MBL consisted of one small building, a few skiffs, and a dock. In 1902 funding for the laboratory was so great that the corporation and board of the MBL considered transferring the laboratory to the
Carnegie Institution of Washington to make the MBL Carnegie’s permanent marine research laboratory. Lillie and Whitman opposed the transfer and convinced the board to reverse its offer to Carnegie. To this day the MBL remains a laboratory relatively free from layers of outside control.

From 1900 to 1942 Lillie worked tirelessly to improve the laboratories and accommodations for the myriad of scientists who descended on Woods Hole during the summer months. Lillie called upon his brother-in-law to help finance the expansion of the MBL and Crane served as president of the corporation from 1904 through 1924. The Crane laboratory at
MBL was named after the man who financed its building. Lillie was instrumental in making the marine laboratory into one of the leading research laboratories in the world. Not only did he serve as president of the corporation from 1925 to 1942 but he also served as Managing Editor of the MBL’s scientific publication
The Biological Bulletin for twenty-five years.

Lillie was an outstanding administrator and teacher but the depth of his research in embryology and development is also remarkable. His early research primarily dealt with
egg cleavage and early development in invertebrates. Although his early cell lineage work was mainly descriptive and comparative, it helped lay the foundation for experimental studies by
Wilhelm Roux in 1888 and
Hans Driesch in 1891. Lillie heavily influenced his former student and noted embryologist
Ernest E. Just to continue working with Nereis to show the relationship of
egg cleavage planes to the entry point of
sperm.

In 1903 and 1904 Lillie published several papers on his studies of the chick embryo. Included in the papers was discussion about the formation of the amnion and his experiments with cauterizing parts of the
embryo to see how further development of parts of the embryo were affected. Lillie had always shown interest in the chick embryo. He was convinced that chick embryos were the best choice for almost any type of experimental work of embryological problems.

From 1910 to 1921 Lillie’s research centered on fertilization in the
annelid Nereis limbata and
sea urchins Arbacia punctulata and Strongylocentrotus franciscanus and S. purpuratus. Lillie proposed that there were specific substances (fertilizin and antifertilizin) secreted by egg and sperm. Part of Lillie’s
“fertilizin theory” likened the interaction between gametes to that of the lock-and-key fashion of antibodies and antigens. This was notable in that Lillie applied the then current immunological concepts and terminology to that of fertilization.

Lillie’s investigation of the factors influencing the development of
freemartins (sterile genetic female calves born as a twin to fertile male calves) helped Lillie answer the question of how sexually indifferent embryos at the beginning of development later turn into males or females. Beginning in 1914 Lillie worked with stockmen in the Swift and Company stockyard to obtain fifty-five pairs of in utero fetal twins from freshly slaughtered pregnant cows. In 1917 Lillie published his study in the
Journal of Experimental Zoology with the finding that freemartin bovine twins are non-identical and that they share the same
placenta, allowing for blood to be freely exchanged between the twin fetuses. A male’s testes form early in development and “masculinizing” substances (hormones) are released and circulate through the fused
umbilical arteries. Lillie concluded that the freemartin was a genetic female calf that had certain male sexual characteristics due to the action of a fetal male sex hormone. This work led to the concept that once a gene directs a gonad to differentiate into a testes or an ovary, accessory reproductive structures in genetic males develop in the male direction due to the presence of male hormones. Genetic females develop rudimentary reproductive structures because they are not inhibited from developing due to a lack of male sex hormones. Lillie’s research with freemartins introduced the notion of the nature and action of sex hormones to embryologists when little was known about hormones. Soon, others at the University of Chicago attempted to produce freemartins in birds (
Benjamin Willier) and mammals (
Carl R. Moore). Castration experiments in fetal mammals in utero and research in the isolation and purification of
sex hormones was undertaken in other laboratories at the University of Chicago. The subsequent work stemming from Lillie’s freemartin investigation helped form the field of reproductive biology.

Even after retiring from the University of Chicago in 1935, Lillie continued with his sex hormone studies by investigating the physiology and development of bird feathers. He used the Brown Leghorn fowl, a bird that displays a notable
sexual dimorphism in feather color and patterns. He collaborated with
Mary Juhn and
His Wang to discover that embryonic feather papillae all start out with the same background color. Further feather coloration and patterns develop in an orderly fashion in response to both female sex hormones and thyroxin. Part of his research involved using castrated males into which injections of estrogens and thyroxin were given to induce “female” feather colorings in the birds’ regenerating feathers.

From 1935 to 1939 Lillie was president of the National Academy of Sciences and in 1935 to 1936 was chairman of the
National Research Council. To date he is the only person ever to have held both leadership positions in the two organizations at the same time. Lillie was also appointed chairman of the National Academy of Sciences Oceanographic Committee to study the financing and construction of an Institute of Oceanography. In 1930 he helped secure a three million dollar grant from the Rockefeller Foundation to help locate and build the Oceanographic Institute next door to the MBL. Lillie served as president of the
Woods Hole Oceanographic Institute from 1930 to 1939.

Lillie died of a stroke on 5 November 1947 in Billings Hospital at the University of Chicago, the campus at which his professional life had so intimately been connected. Thus ended the career of one of the world’s foremost embryologists and science administrators—a career that science historian Philip J. Pauly identified as having helped make biology the first science in which Americans became internationally recognized.

Sources

Lillie, Frank R. The Problems of Fertilization. Chicago: University of Chicago Press, 1919.

Manning, Kenneth J. Black Apollo of Science: The Life of Ernest Everett Just. New York, Oxford University Press, 1983.

Mitman, Gregg. The State of Nature: Ecology, Community, and American Social Thought 1900–1950. Chicago: University of Chicago Press, 1992.

Frank R. Lillie was born in Toronto, Canada, on 27 June 1870. His mother was Emily Ann Rattray and his father was George Waddell Little, an accountant and co-owner of a wholesale drug company. While in high school Lillie took up interests in entomology and paleontology but went to the University of Toronto with the aim of studying ministry. He slowly became disillusioned with this career choice and decided to major in the natural sciences. It was during his senior year that he developed his lifelong interest in embryology.

Jacques Loeb experimented on embryos in Europe and the United States at the end of the nineteenth and beginning of the twentieth centuries. Among the first to study embryos through experimentation, Loeb helped found the new field of
experimental embryology. Notably, Loeb showed scientists how to induce
artificial parthenogenesis, thus refuting the idea that spermatozoa alone were necessary to develop eggs into embryos and confirming the idea that the chemical constitution of embryos’ environment affected their development. Furthermore, Loeb’s work showed that scientists could manipulate materials in a laboratory to create, as he called the process, the beginning stages of life.

Jacques Loeb was born in the Prussian town of Mayen to Barbara and Benedict Loeb in 1859. Named “Isaak,” he changed his name to “Jacques” just prior to entering the
University of Strassburg in 1880. At Strassburg Loeb studied with the physiologist
Friedrich Goltz and there he earned his MD in 1884.

Until 1891 Loeb taught and researched at various institutions, including the
Naples Zoological Station in the winters of 1889 and 1890. In 1890 he met and married
Anne Leonard, an American philologist. Moving to the United States, Loeb taught at
Bryn Mawr College for a year prior to accepting an assistant professorship with the
University of Chicago in 1892. While at Bryn Mawr Loeb met and initially disliked the young
Thomas Hunt Morgan. At Chicago Loeb struggled to get along with
Charles Otis Whitman, and to tolerate what Loeb termed the
“romantic evolutionism” espoused most prominently by Chicago’s
John Dewey.

After a decade at Chicago, Loeb moved to the
University of California at Berkeley for eight years. In California he experimented at Stanford University’s
Hopkins Marine Station in Pacific Grove. He then returned east to New York’s
Rockefeller Institute for Medical Research in 1910, working there until he died in 1924. During his career, he conducted many of his experiments at Bryn Mawr, Chicago, Rockefeller, and at the
Marine Biological Laboratory in Woods Hole, Massachusetts.

Loeb’s early work at Strassburg was on brain physiology. Having judged the field’s dominant theory misguided, he decided to instead study traditional issues in biology such as embryology. He used experiments to answer questions about development and embryology rather than observation alone, which was the common method of biological inquiry. Loeb looked to the scientific practices of physiologists Goltz,
Eduard Pflüger, and the eminent plant physiologist
Julius Sachs for methodological inspiration.

Loeb worked with Sachs extensively while the former worked at the University of Würzburg for two years starting in 1886. Loeb then developed a correspondence and friendship with the Austrian physicist and philosopher
Ernst Mach, whose writings provided the theoretical foundation for Loeb’s work. For Loeb, any explanation of phenomena could come only from manipulating the physical structures of things and their chemical makeup. That conception of scientific knowledge, based on physicochemical manipulations, paralleled
Wilhelm Roux’s mechanistic conception of science encapsulated in
Entwicklungsmechanik.

Like Mach, Loeb thought science was not a mere description of nature but was instead a tool for humans to interact with nature. To him, a biologist was like an engineer and organisms were biologists’ material. Loeb earned his reputation as a biologist-engineer in 1899 when he published experimental results showing
artificial parthenogenesis in
sea urchins, in “
On the Nature of the Process of Fertilization and the Artificial Production of Normal Larvae (Plutei) from the Unfertilized Eggs of the Sea Urchin”.

Artificial parthenogenesis is the human manipulation of egg cells causing embryonic development without spermatozoa. Loeb manipulated unfertilized sea urchin eggs with inorganic solutions of salt water. The result was eggs developing into larvae, or early stage embryos, which he later reproduced using frog eggs. Loeb elaborated on his results in 1913’s
Artificial Parthenogenesis and Fertilization.

Loeb’s 1899 results stirred the popular presses, and many viewed him as a creator of life, perhaps with the ability to engineer new types of organisms. Loeb was never able to create new forms of life, as he intended, but he inspired scientists such as
John Howard Northrop,
John Broadus Watson,
Hermann Joseph Müller,
Burrhus Frederic (B. F.) Skinner, and
Gregory Pincus. Loeb also contributed to the study of
animal tropisms (environment-caused orientation).

Loeb helped transform biology into a largely experimental science. His 1912
The Mechanistic Conception of Life established his reputation as a researcher who treated organisms as machines. In that work, he stated that biologists explain organic phenomena only when they could control those phenomena. Loeb later believed that biologists explain phenomena by detailing the mechanisms, the step-by-step processes, by which a component of an organism achieves its function in physical and chemical terms. In
The Organism as a Whole (1916), Loeb discussed how a mechanist could investigate organisms considered as wholes. His commitment to physicochemical explanations led him to study protein chemistry for the last few years of his life. Loeb died in 1924.

Sources

Loeb, Jacques. “On the Nature of the Process of Fertilization and the Artificial Production of Normal Larvae (Plutei) from the Unfertilized Eggs of Sea Urchins.” The American Journal of Physiology 3 (1899): 135–38.

Loeb, Jacques. The Organism as a Whole: From a Physicochemical Viewpoint. New York: G.P. Putnam’s Sons, 1916.

Jacques Loeb experimented on embryos in Europe and the United States at the end of the nineteenth and beginning of the twentieth centuries. Among the first to study embryos through experimentation, Loeb helped found the new field of experimental embryology. Notably, Loeb showed scientists how to create artificial parthenogenesis, thus refuting the idea that spermatozoa alone were necessary to develop eggs into embryos and confirming the idea that the chemical constitution of embryos environment affected their development.